Nesting Migrations and Reproductive Biology of the Mona Rhinoceros Iguana, Cyclura Stejnegeri
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Herpetological Conservation and Biology 11(Monograph 6):197–213. Submitted: 2 September 2014; Accepted: 12 November 2015; Published: 12 June 2016. NESTING MIGRATIONS AND REPRODUCTIVE BIOLOGY OF THE MONA RHINOCEROS IGUANA, CYCLURA STEJNEGERI 1,3 1 2 NÉSTOR PÉREZ-BUITRAGO , ALBERTO M. SABAT , AND W. OWEN MCMILLAN 1Department of Biology, University of Puerto Rico – Río Piedras, San Juan, Puerto Rico 00931 2Smithsonian Tropical Research Institute, Apartado 0843-03092 Panamá, República of Panamá 3 Current Address: Universidad Nacional de Colombia, Sede Orinoquía, Grupo de Investigación en Ciencias de la Orinoquía (GICO), km 9 vía Tame, Arauca, Colombia 3Corresponding author, e-mail: [email protected] Abstract.—We studied the nesting migrations and reproductive ecology of the endangered Mona Rhinoceros Iguana Cyclura stejnegeri at three localities from 2003 to 2006. Female movements while seeking a nesting site ranged from 0.3 to 12.8 km, were mostly erratic. Time elapsed between mating and oviposition averaged 30 ± 5 days, while the nesting period lasted four weeks (July to early August). Nest site fidelity by females in consecutive years was 50%, although non-resident females at one study site used the same beach 72% of the time. Clutch size averaged 14 eggs and was positively correlated with female snout-vent length (SVL). Egg length was the only egg size variable correlated negatively with female size. Incubation temperatures averaged 32.8° C (2005) and 30.2° C (2006) and fluctuated up to 9° C. Overall hatching success from 2003–2005 was 75.9%. Some nests failed as a result of flooding of the nest chamber and in one case a nest was destroyed by feral pigs. Hatchling sex ratio was close to 1:1 for all individual nests and all hatchlings collected throughout the study. Hatchling morphometrics (SVL, body mass) were not related to female size. Egg predation by pigs was low compared to previous reports from the 1970s, which reached levels of up to 100% in some years. The dramatic increase in hatching success may be the result of fencing the most important iguana nesting areas in 1985, an initiative that is maintained until now, to prevent feral pig incursions. Resumen.—Del 2003 al 2006 estudiamos la migraciones asociadas a la búsqueda de sitios para la anidación y la ecología reproductiva de la iguana Cyclura stejnegeri en tres localidades de Isla de Mona. Las distancias recorridas por las hembras durante la búsqueda de sitios para anidar fluctuaron entre 0.3 y 12.8 km con desplazamientos en su mayoría erráticos. El tiempo promedio entre el apareamiento y la ovoposición fue de 30 ± 5 días y la época de anidación duró cuatro semanas entre julio y agosto. Las hembras mostraron fidelidad al sitio de anidaje en un 50% de los casos, aunque el 72% de hembras no residentes en una playa retornaron a la misma en años consecutivos. El promedio de huevos por nidada fue de 14 y estuvo positivamente correlacionado con el tamaño (longitud hocico-cloaca, SVL) de la hembra. De las dimensiones de los huevos, solo el largo del huevo estuvo negativamente correlacionado con el tamaño de la hembra. Las temperaturas de incubación promedio fueron de 32.8° C (2005) y 30.2° C (2006) con fluctuaciones hasta de 9° C. Algunos nidos fueron destruidos debido a las fuertes lluvias que inundaron la cámara del nido y solo un nido fue destruido por cerdos silvestres. La proporción sexual de los neonatos fue de 1:1 para los nidos individuales y para todos los neonatos colectados en el estudio. La morfometría de los neonatos (SVL, peso) no estuvo relacionada con el tamaño de la madre. La depredación de nidos por cerdos fue baja comparada con los años setenta cuando podía alcanzar valores hasta del 100% en algunos años. El incremento sustancial en el éxito de eclosión puede ser el resultado de la instalación de cercas en 1985 en algunas áreas de anidaje para evitar la depredación de nidos por parte de cerdos silvestres. Key Words.—Caribbean Iguanine; lizard; nesting ecology; reproductive success; rock iguana INTRODUCTION (Alberts 2007; Pérez-Buitrago et al. 2008; Burton and Rivera-Milán 2014), translocations (Knapp 2000, 2001; Caribbean Ground Rock Iguanas (11 species, eight Knapp and Hudson 2004; Wilson et al. 2004), and subspecies; ITWG this volume) of the genus Cyclura are eradication/control of feral animal species (Mitchell et among the most endangered lizards in the world (Alberts al. 2002; Donlan et al. 2003; Campbell et al. 2004; 2000). Factors affecting most of the populations of these Hayes et al. 2004; Gerber 2007). However, knowledge species include habitat degradation, illegal pet trade, and of Cyclura biology is relatively poor due to the negative interactions with exotic animals (Alberts 2000). remoteness of islands on which some species live, and/or Current actions to augment some Cyclura populations the lack of financial/logistic resources to study these include captive breeding and headstarting initiatives long-lived lizards (Iverson et al. 2004). Copyright © 2016. Néstor Pérez-Buitrago; All Rights Reserved. Pérez-Buitrago, Néstor, AlBerto M. SaBat, and W. Owen McMillan. 2016. Nesting migrations and reproductive Biology of the Mona Rhinoceros Iguana, Cyclura stejnegeri. Pp. 197–213 In Iguanas: Biology, Systematics, and Conservation. Iverson, J.B., T.D. Grant, C.R. Knapp, and S.A. Pasachnik (Eds.). Herpetological Conservation and Biology 11(Monograph 6). Pérez-Buitrago et al.— Reproductive Biology of the Mona Rhinoceros Iguana. Knowledge of the reproductive biology of the genus However, a large part of the coastal plain was planted Cyclura is a critical component to the evaluation of their with exotic species such as Casuarina equisetifolia and population trends, but studies addressing their Swietenia mahogany (Diaz 1984; Cintrón and Rogers reproduction have been limited to descriptions of basic 1991), thus reducing the availability of appropriate and aspects such as timing of nesting and hatching, clutch contiguous nesting sites. Nonetheless, it still constitutes size, egg and hatchling dimensions, and hatching success an estimated 74% of all nesting habitat available on the based on relatively small sample sizes and over short island. Currently, there are small sunny and sandy areas periods of time (but see Iverson et al. 2004; Knapp et al. that apparently have not been affected by human activity 2006). This information has been used to analyze life- recently, as well as areas that were cleared by the DRNA- history trait patterns among species/populations PR in 1992 to provide additional suitable zones for (Wiewandt 1982; Iverson et al. 2004), but its utility in nesting females. In addition to the nesting areas located in developing population viability models that may be used the southwestern coastal plain, there are a few narrow (< to guide specific conservation actions is limited. 150 m wide) beaches available that represent the third Moreover, there are still several aspects of reproduction type of open area available for iguana nesting. in Cyclura that are poorly documented, such as details of This study was conducted at three locations of Mona their nesting migrations. From a conservation Island representing two of the three nesting habitats perspective, knowledge of the factors that may be described above. The Lighthouse site is located near the reducing the availability of nesting areas and/or reducing eastern coast of Mona Island on its limestone plateau, hatching success rates is critical. lacks human disturbance and does not contain well- In this study we document the reproductive biology of defined nesting areas (Fig. 1). The second area, the Mona Rhinoceros Iguana (Cyclura stejnegeri) using Sardinera Beach, is located in the southwestern coastal radio-telemetry and mark-recapture techniques. During plain near the site of the DRNA-PR facilities, and is a three reproductive seasons (2003–2005), we documented highly disturbed area close to the zones in which the nest timing, clutch size, egg dimensions, hatchling size, native vegetation was replaced by exotic tree species. reproductive effort, and the nesting migrations The third study site, Pájaros Beach, is a narrow but undertaken by females to reach nesting sites. In elongated beach in the southeast where the predominant addition, we compared the spatial variation in clutch size vegetation type is “cliff forest” (Cintrón and Rogers and hatching success across study sites. We also 1991). This area has a small camping facility used by evaluate the effect of management actions taken by the tourists approximately 30% of the year, and contains Department of Natural Resources of Puerto Rico some open zones appropriate for nesting. (DRNA-PR) in 1982 on reproductive success. These actions included fencing some nesting areas to prevent Field data collection.—From 2003 to 2005 we feral pig incursions and creating areas free of vegetation captured female iguanas at the three study sites using to increase available nesting habitat. nets. Most capture effort was concentrated during April, June–July, and October–November. For each captured MATERIALS AND METHODS iguana, we recorded the snout-vent length (SVL) to the nearest 1 mm, body mass (BM) to the nearest 0.1 kg, tail Study site.—Mona Island is located in the middle of length (TL) and tail breaks to the nearest 1 mm. If sex the Mona channel, between the Dominican Republic and was externally unclear, we determined it by probing Puerto Rico. It is an oceanic island with a subtropical (Schaeffer 1934; Dellinger and Von Hegel 1990). We dry forest climate (Ewel and Whitmore 1973). Most of marked iguanas externally with a unique combination of the island’s perimeter is characterized by vertical cliffs color beads attached to the dorsal crest (Rodda et al. 45 m in height that also delimit the largest habitat type of 1988) and internally with a passive integrated the island (93%), the relatively undisturbed limestone transponder (PIT) tag (AVID®).